Kurimwa kwemupunga kuri kudzikira nekuda kwekushanduka kwemamiriro ekunze uye kusiyana kwemamiriro ekunze muColombia.Vatongi vekukura kwezvirimwazvakashandiswa senzira yekuderedza kushushikana kwekupisa muzvirimwa zvakasiyana-siyana. Nokudaro, chinangwa chechidzidzo ichi chaive chekuongorora mhedzisiro yemuviri (kutenderera kwemupunga, kutenderera kwemupunga, huwandu hwechlorophyll, chiyero cheFv/Fm chemhando mbiri dzemupunga dzekutengesa dzakabatwa nekupisa kwakabatana (kupisa kwakanyanya masikati neusiku), tembiricha yedenga uye huwandu hwemvura) uye biochemical variables (malondialdehyde (MDA) uye huwandu hweprolinic acid). Kuedza kwekutanga nekwechipiri kwakaitwa uchishandisa zvirimwa zvemhando mbiri dzemupunga Federrose 67 ("F67") naFederrose 2000 ("F2000"), zvichiteerana. Kuedza kwese kwakaongororwa pamwe chete seongororo dzakatevedzana. Kurapa kwakasimbiswa kwaive kunotevera: absolute control (AC) (zvirimwa zvemupunga zvakarimwa patembiricha yepamusoro (tembiricha yemasikati/husiku 30/25°C)), heat stress control (SC) [zvirimwa zvemupunga zvakabatwa nekupisa kwakabatana chete (40/ 25°C). 30°C)], uye zvirimwa zvemupunga zvakamanikidzwa uye zvakapfapfaidzwa nezvinogadzirisa kukura kwezvirimwa (stress+AUX, stress+BR, stress+CK kana stress+GA) kaviri (mazuva mashanu zvisati zvaitika uye mazuva mashanu mushure mekumanikidzwa kupisa). Kupfapfaidza neSA kwakawedzera huwandu hwese hwechlorophyll hwemarudzi ese ari maviri (huremu hutsva hwezvirimwa zvemupunga "F67″ uye "F2000″ hwaive 3.25 uye 3.65 mg/g, zvichiteerana) zvichienzaniswa nezvirimwa zveSC (huremu hutsva hwezvirimwa zve "F67″ hwaive 2.36 uye 2.56 mg). g-1)" uye mupunga "F2000", kushandiswa kweCK pamashizha kwakavandudzawo kufambisa kwemupunga "F2000″ zvirimwa (499.25 vs. 150.60 mmol m-2 s) zvichienzaniswa nekudzora kupisa. Kumanikidzwa kupisa, tembiricha yekorona yemuti inoderera ne2–3 °C, uye huwandu hweMDA muzvirimwa hunoderera. Ruzivo rwekushivirira runoratidza kuti kushandiswa kweCK (97.69%) neBR (60.73%) pamashizha kunogona kubatsira kuderedza dambudziko rekupisa kwakabatana. kunyanya muzvirimwa zvemupunga zveF2000. Mukupedzisa, kupfapfaidza BR kana CK pamashizha kunogona kutariswa senzira yekubatsira kuderedza migumisiro yakaipa yekupisa kwakabatana pamaitiro emuviri wezvirimwa zvemupunga.
Mupunga (Oryza sativa) ndewemhuri yePoaceae uye ndeimwe yezviyo zvinorimwa zvakanyanya pasi rose pamwe chete nechibage negorosi (Bajaj naMohanty, 2005). Nzvimbo iri kurimwa mupunga i617,934 hectares, uye kugadzirwa kwenyika yose muna 2020 kwaive matani 2,937,840 neavhareji yegoho rematani 5.02/ha (Federarroz (Federación Nacional de Arroceros), 2021).
Kudziya kwepasi rose kuri kukanganisa zvirimwa zvemupunga, zvichikonzera mhando dzakasiyana dzematambudziko asina kugayiwa dzakadai sekupisa kwakanyanya uye nguva dzekusanaya kwemvura. Kuchinja kwemamiriro ekunze kuri kukonzera kukwira kwekupisa kwepasi rose; Tembiricha dziri kutarisirwa kukwira ne1.0–3.7°C muzana remakore rechi21, izvo zvinogona kuwedzera kuwanda kwekupisa. Kukwira kwekupisa kwezvakatipoteredza kwakanganisa mupunga, zvichikonzera kuti goho rezvirimwa riderere ne6–7%. Kune rumwe rutivi, kuchinja kwemamiriro ekunze kunotungamirawo kumamiriro ekunze asina kunaka kuzvirimwa, senge nguva dzekusanaya kwemvura kwakanyanya kana tembiricha dzakakwirira munzvimbo dzinopisa nedzinodziya. Pamusoro pezvo, zviitiko zvakasiyana-siyana zvakaita seEl Niño zvinogona kutungamira kukushomeka kwekupisa uye kuwedzera kukuvara kwezvirimwa mune dzimwe nzvimbo dzinopisa. MuColombia, tembiricha munzvimbo dzinogadzira mupunga dziri kutarisirwa kuwedzera ne2–2.5°C panosvika 2050, zvichideredza kugadzirwa kwemupunga uye kukanganisa kuyerera kwezvigadzirwa kumisika uye cheni dzezviwanikwa.
Zvirimwa zvakawanda zvemupunga zvinorimwa munzvimbo dzine tembiricha dziri pedyo nedzemhando yepamusoro yekukura kwezvirimwa (Shah et al., 2011). Zvakataurwa kuti tembiricha yepamusoro yemasikati nehusiku yekukura nekukura kwemupungakazhinji kacho 28°C uye 22°C, zvichiteerana (Kilasi et al., 2018; Calderón-Páez et al., 2021). Kupisa kuri pamusoro peizvi kunogona kukonzera nguva dzekupisa zvine mwero kusvika zvakanyanya panguva yekukura kwemupunga (kurima, kunhuhwirira, kutumbuka, uye kuzadza zviyo), nokudaro zvichikanganisa goho rezviyo. Kudzikira kwegoho uku kunonyanya kukonzerwa nenguva refu yekudziya, iyo inokanganisa mashandiro emiti. Nekuda kwekudyidzana kwezvinhu zvakasiyana-siyana, zvakaita senguva yekushushikana uye tembiricha yepamusoro inosvika, kupisa kunogona kukonzera kukuvara kusingadzoreki kumetabolism yemiti nekukura kwayo.
Kushushikana nekupisa kunokanganisa maitiro akasiyana-siyana emuviri uye emakemikari mumiti. Kuchinjana kwemashizha ndeimwe yemaitiro anonyanya kukanganiswa nekupisa mumiti yemupunga, sezvo huwandu hwefotosinthesiti hunoderera ne50% kana tembiricha dzezuva nezuva dzichipfuura 35°C. Mhinduro dzemuviri dzemiti yemupunga dzinosiyana zvichienderana nerudzi rwekupisa. Semuenzaniso, mwero wephotosynthesis uye kufambisa kwedumbu zvinodziviswa kana zvirimwa zvikasangana nekupisa kwakanyanya masikati (33–40°C) kana tembiricha yepamusoro masikati neusiku (35–40°C masikati, 28–30°C). C zvinoreva husiku) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Kupisa kwakanyanya husiku (30°C) kunokonzera kudzikiswa kwefotosinthesiti asi kunowedzera kufema kweusiku (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Pasinei nenguva yekushushikana, kushushikana kwekupisa kunokanganisawo huwandu hwe chlorophyll mumashizha, chiyero che chlorophyll variable fluorescence kusvika ku chlorophyll fluorescence yakanyanya (Fv/Fm), uye Rubisco activation mumiti yemupunga (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso et al., 2014).
Kuchinja kwemakemikari emuviri chimwe chinhu chinoitwa nemiti kuti ikwanise kugadzikana nekupisa (Wahid et al., 2007). Kuwanda kweproline kwave kuchishandiswa sechiratidzo chebiochemical chekumanikidzwa kwemiti (Ahmed naHassan 2011). Proline inoita basa rakakosha mukugadzirwa kwemiti sezvo ichishanda senzvimbo ine kabhoni kana nitrogen uye senzira yekusimbisa membrane pasi pemamiriro ekupisa kwakanyanya (Sánchez-Reinoso et al., 2014). Kupisa kwakanyanya kunokanganisawo kugadzikana kwemembrane kuburikidza ne lipid peroxidation, zvichikonzera kuumbwa kwe malondialdehyde (MDA) (Wahid et al., 2007). Nokudaro, huwandu hweMDA hwakashandiswawo kunzwisisa hunhu hwema membrane emasero pasi pekupisa (Cao et al., 2009; Chavez-Arias et al., 2018). Pakupedzisira, kumanikidzwa kwekupisa kwakabatana [37/30°C (masikati/husiku)] kwakawedzera huwandu hwekubuda kwe electrolyte uye huwandu hwe malondialdehyde mupunga (Liu et al., 2013).
Kushandiswa kwezvinodzora kukura kwezvirimwa (GRs) kwakaongororwa kuderedza migumisiro yakaipa yekumanikidzwa nekupisa, sezvo zvinhu izvi zvichibatanidzwa zvakanyanya mukupindura kwezvirimwa kana nzira dzekudzivirira muviri kubva mukumanikidzwa kwakadaro (Peleg naBlumald, 2011; Yin et al. et al., 2011; Ahmed et al., 2015). Kushandiswa kwezviwanikwa zvemajini kunze kwenyika kwave nemhedzisiro yakanaka pakushivirira kupisa muzvirimwa zvakasiyana-siyana. Zvidzidzo zvakaratidza kuti ma phytohormones akadai se gibberellins (GA), cytokinins (CK), auxins (AUX) kana brassinosteroids (BR) anotungamira mukuwedzera kwezvimiro zvakasiyana-siyana zvemuviri uye zvebiochemical (Peleg naBlumald, 2011; Yin et al. Ren, 2011; Mitler et al., 2012; Zhou et al., 2014). MuColombia, kushandiswa kwezviwanikwa zvemajini kunze kwenyika uye kukanganisa kwazvinoita pazvirimwa zvemupunga hazvina kunyatsonzwisiswa uye kudzidzwa. Zvisinei, ongororo yapfuura yakaratidza kuti kupfapfaidza BR pamashizha kunogona kuvandudza kushivirira mupunga nekuvandudza hunhu hwekutsinhana kwegasi, chlorophyll kana huwandu hweproline mumashizha emhodzi dzemupunga (Quintero-Calderón et al., 2021).
MaCytokinins anobatsira pakuita kuti zvirimwa zvisanyanya kunetseka, kusanganisira kupisa (Ha et al., 2012). Pamusoro pezvo, zvakanzi kushandiswa kweCK kunze kunogona kuderedza kukuvara kwekupisa. Semuenzaniso, kushandiswa kwezeatin kunze kwakawedzera mwero we photosynthetic, chlorophyll a uye b zviri mukati, uye kushanda zvakanaka kwekufambisa maerekitironi mu creeping bentgrass (Agrotis estolonifera) panguva yekupisa (Xu naHuang, 2009; Jespersen naHuang, 2015). Kushandiswa kwezeatin kunze kunogonawo kuvandudza mashandiro eantioxidant, kuwedzera kugadzirwa kwemapuroteni akasiyana-siyana, kuderedza kukuvara kwe reactive oxygen (ROS) uye malondialdehyde (MDA) mumatishu emiti (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar et al., 2020).
Kushandiswa kwegibberellic acid kwakaratidzawo mhinduro yakanaka pakumanikidzwa nekupisa. Zvidzidzo zvakaratidza kuti GA biosynthesis inobatanidza nzira dzakasiyana dzekugadzirisa muviri uye inowedzera kushivirira mumamiriro ekupisa kwakanyanya (Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi et al. (2020) vakaona kuti kupfapfaidza GA yekunze (25 kana 50 mg*L) pamashizha kunogona kuwedzera mwero we photosynthesis uye antioxidant mumiti yeorenji ine kupisa kwakanyanya zvichienzaniswa nemiti inotonga. Zvakaonekwawo kuti kushandiswa kweHA kunze kunowedzera huwandu hwehunyoro, chlorophyll uye carotenoid uye kunoderedza lipid peroxidation mumuti we date palm (Phoenix dactylifera) uri pasi pekupisa kwakanyanya (Khan et al., 2020). Auxin inoitawo basa rakakosha mukudzora mhinduro dzekukura dzinochinjika kune mamiriro ekupisa kwakanyanya (Sun et al., 2012; Wang et al., 2016). Uyu mugadziri wekukura unoshanda sechiratidzo chebiochemical mumabasa akasiyana-siyana akadai sekugadzirwa kweproline kana kuora pasi pekumanikidzwa kweabiotic (Ali et al. 2007). Pamusoro pezvo, AUX inowedzerawo basa reantioxidant, izvo zvinotungamira kudzikira kweMDA muzvirimwa nekuda kwekudzikira kwemafuta emudumbu (Bielach et al., 2017). Sergeev et al. (2018) vakaona kuti muzvirimwa zvepizi (Pisum sativum) pasi pekumanikidzwa kwekupisa, huwandu hweproline - dimethylaminoethoxycarbonylmethyl)naphthylchloromethyl ether (TA-14) hunowedzera. Mukuyedza kumwe chete, vakaonawo huwandu hwakaderera hweMDA muzvirimwa zvakarapwa zvichienzaniswa nezvirimwa zvisina kurapwa neAUX.
Brassinosteroids ndeimwe mhando yezvinogadzirisa kukura kwezvirimwa zvinoshandiswa kuderedza mhedzisiro yekumanikidzwa nekupisa. Ogweno et al. (2008) vakataura kuti exogenous BR spray yakawedzera net photosynthetic rate, stomatal conductance uye maximum rate yeRubisco carboxylation yematomatisi (Solanum lycopersicum) zvirimwa zviri pasi pekupisa kwemazuva masere. Kupfapfaidza epibrassinosteroids nefoliar kunogona kuwedzera net photosynthetic rate yezvirimwa zvecucumber (Cucumis sativus) zviri pasi pekupisa (Yu et al., 2004). Pamusoro pezvo, kushandiswa kweBR kunze kunonotsa kuora kwechlorophyll uye kunowedzera kushanda zvakanaka kwemvura uye goho guru rePSII photochemistry muzvirimwa zviri pasi pekupisa (Holá et al., 2010; Toussagunpanit et al., 2015).
Nekuda kwekushanduka kwemamiriro ekunze uye kushanduka, zvirimwa zvemupunga zvinotarisana nenguva dzekupisa kwemazuva ese (Lesk et al., 2016; Garcés, 2020; Federarroz (Federación Nacional de Arroceros), 2021). Mumiti inonzi phenotyping, kushandiswa kwe phytonutrients kana biostimulants kwakadzidzwa senzira yekuderedza kupisa kwemhepo munzvimbo dzinorimwa mupunga (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Pamusoro pezvo, kushandiswa kwezvinhu zvinochinja-chinja zvemakemikari nemuviri (tembiricha yemashizha, stomatal conductance, chlorophyll fluorescence parameters, chlorophyll uye relative water content, malondialdehyde uye proline synthesis) chishandiso chakavimbika chekuongorora zvirimwa zvemupunga zviri pasi pekupisa kwakanyanya munharaunda nepasi rese (Sánchez -Reynoso et al., 2014; Alvarado-Sanabria et al., 2017; Zvisinei, tsvakiridzo pamusoro pekushandiswa kwefoliar phytohormonal sprays mumupunga padanho remunharaunda ichiri shoma. Nokudaro, kudzidza kwemaitiro emuviri nemakemikari ekushandiswa kwezvinodzora kukura kwezvirimwa kwakakosha zvikuru pakupa mazano ezvekurima anoshanda eizvi. kugadzirisa migumisiro yakaipa yenguva yekushushikana kwakanyanya kwekupisa mumupunga. Nokudaro, chinangwa chechidzidzo ichi chaive chekuongorora physiological (stomatal conductance, chlorophyll fluorescence parameters uye relative water content) uye mhedzisiro yebiochemical yekushandiswa kwefoliar kwezvinodzora kukura kwezvirimwa zvina (AUX, CK, GA uye BR). (Photosynthetic pigments, malondialdehyde uye proline contents) Mhando mbiri dzemupunga wekutengesa dzakasangana nekupisa kwakanyanya (kupisa kwakanyanya masikati/husiku).
Muchidzidzo ichi, kuyedza kwakaitwa kaviri kwakazvimirira. Marudzi eFederrose 67 (F67: rudzi rwegenotype rwakakura mukupisa kwakanyanya mumakore gumi apfuura) uye Federrose 2000 (F2000: rudzi rwegenotype rwakakura mumakore gumi ekupedzisira ezana remakore rechi20 rwuchiratidza kusabatwa nehutachiona hwemashizha machena) akashandiswa kekutanga. mbeu. uye kuyedza kwechipiri, zvichiteerana. Marudzi ese ari maviri anorimwa zvakanyanya nevarimi vekuColombia. Mbeu dzakadyarwa mumatireyi e10-L (kureba 39.6 cm, upamhi 28.8 cm, kureba 16.8 cm) dzine ivhu rejecha rine 2% organic matter. Mbeu shanu dzakamera kare dzakadyarwa mutireyi yega yega. Mapallet akaiswa mugreenhouse yeFaculty of Agricultural Sciences yeNational University of Colombia, Bogotá campus (43°50′56″ N, 74°04′051″ W), pakukwirira kwe2556 m pamusoro pegungwa (asl). m.) uye zvakaitwa kubva muna Gumiguru kusvika Zvita 2019. Kuedza kumwe chete (Federroz 67) uye kuedza kwechipiri (Federroz 2000) mumwaka mumwe chete wa2020.
Mamiriro ekunze ari mugreenhouse panguva yega yega yekudyara ndeaya anotevera: tembiricha yemasikati nehusiku 30/25°C, hunyoro hwe60~80%, nguva yekutora mifananidzo yechisikigo maawa gumi nemaviri (mwaranzi inoshanda nephotosynthetically 1500 µmol (photons) m-2 s-). 1 masikati). Zvirimwa zvakaiswa fetereza zvichienderana nehuwandu hwechinhu chimwe nechimwe mazuva makumi maviri mushure mekubuda kwembeu (DAE), sekureva kwaSánchez-Reinoso et al. (2019): 670 mg nitrogen pachirimwa, 110 mg phosphorus pachirimwa, 350 mg potassium pachirimwa, 68 mg calcium pachirimwa, 20 mg magnesium pachirimwa, 20 mg sulfur pachirimwa, 17 mg silicon pachirimwa. Zvirimwa izvi zvine 10 mg boron pachirimwa, 17 mg copper pachirimwa, uye 44 mg zinc pachirimwa. Zvirimwa zvemupunga zvakachengetwa kusvika ku47 DAE mukuyedza kwega kwega pazvakasvika padanho re phenological V5 munguva iyi. Zvidzidzo zvekare zvakaratidza kuti danho iri re phenological inguva yakakodzera yekuita zvidzidzo zvekupisa mumupunga (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Mukuyedza kwega kwega, mashandisirwo maviri akasiyana emuchina wekutonga kukura kwemashizha akaitwa. Seti yekutanga yemishonga yekubvisa maruva inonzi foliar phytohormone yakashandiswa mazuva mashanu mushonga wekupisa usati watanga (42 DAE) kugadzirira zvirimwa kuti zvisanyanya kuoma. Imwe mishonga yekubvisa maruva yakapihwa mazuva mashanu mushure mekunge zvirimwa zvasangana nemamiriro ekumanikidzwa (52 DAE). Ma phytohormone mana akashandiswa uye hunhu hwechinhu chimwe nechimwe chinoshanda chakapfapfaidzwa muchidzidzo ichi hwakanyorwa muSupplementary Table 1. Kuwanda kwemishonga yekuderedza kukura kwemashizha yakashandiswa kwaive seizvi: (i) Auxin (1-naphthylacetic acid: NAA) pahuwandu hwe5 × 10−5 M (ii) 5 × 10–5 M gibberellin (gibberellic acid: NAA); GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. Kuwanda uku kwakasarudzwa nekuti kunokonzera mhinduro dzakanaka uye kunowedzera kusasimba kwemiti pakupisa (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Miriwo yemupunga isina spray inodzora kukura kwemiti yakapiswa nemvura yakasanganiswa chete. Miriwo yese yemupunga yakapfapfaidzwa neruoko. Isa 20 ml H2O pamuti kuti unyorovese pamusoro nepazasi pemashizha. Spray dzese dzemashizha dzakashandisa agricultural adjuvant (Agrotin, Bayer CropScience, Colombia) pa0.1% (v/v). Daro riri pakati pehari nesprayer i30 cm.
Kurapa kwekumanikidzwa nekupisa kwakaitwa mazuva mashanu mushure mekupfapfaidza kwemashizha ekutanga (47 DAE) mukuyedza kwega kwega. Zvirimwa zvemupunga zvakatamiswa kubva mugreenhouse kuenda kuimba yekukura ye294 L (MLR-351H, Sanyo, IL, USA) kuti pave nekumanikidzwa nekupisa kana kuchengetedza mamiriro akafanana ezvakatipoteredza (47 DAE). Kurapa kwekumanikidzwa nekupisa kwakaitwa nekuisa imba patembiricha inotevera yemasikati/husiku: tembiricha yepamusoro yemasikati [40°C kwemaawa mashanu (kubva 11:00 kusvika 16:00)] uye nguva yehusiku [30°C kwemaawa mashanu]. Mazuva masere akatevedzana (kubva 19:00 kusvika 24:00). Tembiricha yekumanikidzwa nenguva yekuratidzwa zvakasarudzwa zvichibva pane zvidzidzo zvekare (Sánchez-Reynoso et al. 2014; Alvarado-Sanabía et al. 2017). Kune rumwe rutivi, boka rezvirimwa zvakatamisirwa muimba yekukura zvakachengetwa mugreenhouse patembiricha imwechete (30°C masikati/25°C husiku) kwemazuva masere akatevedzana.
Pakupera kwekuyedza, mapoka ekurapa anotevera akawanikwa: (i) mamiriro ekupisa kwekukura + kushandiswa kwemvura yakasanganiswa [Absolute control (AC)], (ii) mamiriro ekupisa + kushandiswa kwemvura yakasanganiswa [Heat stress control (SC)], (iii) mamiriro ekupisa + auxin application (AUX), (iv) mamiriro ekupisa + gibberellin application (GA), (v) mamiriro ekupisa + cytokinin application (CK), uye (vi) mamiriro ekupisa + brassinosteroid (BR) Appendix. Mapoka aya ekurapa akashandiswa kune ma genotype maviri (F67 naF2000). Kurapa kwese kwakaitwa nenzira isina kurongeka ine makopi mashanu, imwe neimwe ine chirimwa chimwe. Chirimwa chega chega chakashandiswa kuverenga ma variables akasarudzwa pakupera kwekuyedza. Kuyedza kwacho kwakatora 55 DAE.
Kuyerera kwemudumbu (gs) kwakayerwa uchishandisa portable porosometer (SC-1, METER Group Inc., USA) inotangira pa0 kusvika 1000 mmol m-2 s-1, ine mukana wekamuri remuenzaniso we6.35 mm. Kuyera kunotorwa nekubatanidza stomameter probe pashizha rakakura nemuti mukuru wakakura zvizere. Pakurapa kwega kwega, kuverenga kwegs kwakatorwa pamashizha matatu emuti wega wega pakati pa11:00 na16:00 uye kwakayerwa paavhareji.
RWC yakaongororwa zvichienderana nenzira yakatsanangurwa naGhoulam et al. (2002). Pepa rakanyatso shandiswa kuona g rakashandiswawo kuyera RWC. Huremu hutsva (FW) hwakaongororwa pakarepo mushure mekukohwa uchishandisa chikero chedhijitari. Mashizha acho akabva aiswa mumudziyo wepurasitiki wakazadzwa nemvura ndokusiyiwa murima patembiricha yemumba (22°C) kwemaawa makumi mana nemasere. Wobva wayera pachikero chedhijitari wonyora huremu hwakakura (TW). Mashizha akazvimba akaomeswa muchoto pa75°C kwemaawa makumi mana nemasere uye huremu hwawo hwakaoma (DW) hwakanyorwa.
Huwandu hwechlorophyll hwakaongororwa uchishandisa chlorophyll meter (atLeafmeter, FT Green LLC, USA) uye hwakaratidzwa mu atLeaf units (Dey et al., 2016). Kuverengwa kwePSII maximum quantum efficiency (Fv/Fm ratio) kwakanyorwa uchishandisa continuous excitation chlorophyll fluorimeter (Handy PEA, Hansatech Instruments, UK). Mashizha akagadziriswa nerima achishandisa mashizha emashizha kwemaminitsi makumi maviri Fv/Fm isati yaongororwa (Restrepo-Diaz naGarces-Varon, 2013). Mushure mekunge mashizha asviba, baseline (F0) uye maximum fluorescence (Fm) zvakayerwa. Kubva padata iri, variable fluorescence (Fv = Fm – F0), chiyero chevariable fluorescence kusvika pamanicure fluorescence (Fv/Fm), maximum quantum yield yePSII photochemistry (Fv/F0) uye chiyero cheFm/F0 zvakaverengerwa (Baker, 2008; Lee et al. ., 2017). Kuverengwa kwe chlorophyll ne chlorophyll fluorescence kwakatorwa pamashizha akafanana akashandiswa pakuyera gs.
Inenge 800 mg yehuremu hwemashizha matsva yakaunganidzwa sezvinhu zvinoshandura biochemical. Mienzaniso yemashizha yakazosanganiswa mu liquid nitrogen ndokuchengetwa kuti iongororwe zvakanyanya. Nzira ye spectrometric inoshandiswa kufungidzira huwandu hwe chlorophyll a, b uye carotenoid mumatishu yakavakirwa panzira uye ma equation akatsanangurwa naWellburn (1994). Mienzaniso yematishu emashizha (30 mg) yakaunganidzwa uye yakasanganiswa mu 3 ml ye 80% acetone. Mienzaniso yacho yakazoiswa centrifuge (model 420101, Becton Dickinson Primary Care Diagnostics, USA) pa 5000 rpm kwemaminitsi gumi kuti ibvise zvidimbu. Supernatant yakasanganiswa kusvika pahuwandu hwekupedzisira hwe 6 ml nekuwedzera 80% acetone (Sims naGamon, 2002). Huwandu hwechlorophyll hwakaonekwa pa663 (chlorophyll a) uye 646 (chlorophyll b) nm, uye carotenoids pa470 nm uchishandisa spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Nzira ye thiobarbituric acid (TBA) yakatsanangurwa naHodges et al. (1999) yakashandiswa kuongorora membrane lipid peroxidation (MDA). Inenge 0.3 g yemashizha emashizha yakagadziridzwawo mu liquid nitrogen. Samples dzakaiswa pakati pemvura nemhepo pa5000 rpm uye absorbance yakayerwa pa spectrophotometer pa440, 532 uye 600 nm. Pakupedzisira, huwandu hweMDA hwakaverengerwa uchishandisa extinction coefficient (157 M mL−1).
Kuwanda kweproline mumishonga yese kwakaongororwa uchishandisa nzira yakatsanangurwa naBates et al. (1973). Wedzera 10 ml ye3% aqueous solution yesulfosalicylic acid kumushonga wakachengetedzwa wobva waisa sefa kuburikidza neWhatman filter paper (Nhamba 2). Zvadaro 2 ml yefiltrate iyi yakasanganiswa ne2 ml yeninhydric acid uye 2 ml yeglacial acetic acid. Musanganiswa wacho wakaiswa mumvura yekugezera pa90°C kweawa imwe chete. Mira kuita izvi nekuzviisa muchando. Zunza chubhu nesimba uchishandisa vortex shaker wonyungudutsa mhinduro yacho mu4 ml yetoluene. Kuverengwa kweAbsorbance kwakaongororwa pa520 nm uchishandisa spectrophotometer imwechete inoshandiswa pakuenzanisa pigments dzephotosynthetic (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Nzira yakatsanangurwa naGerhards et al. (2016) yekuverenga tembiricha yedenga uye CSI. Mifananidzo yekupisa yakatorwa nekamera yeFLIR 2 (FLIR Systems Inc., Boston, MA, USA) ine kurongeka kwe ±2°C pakupera kwenguva yekushushikana. Isa nzvimbo chena kumashure kwechirimwa kuti utore mifananidzo. Zvakare, mafekitori maviri akatorwa semamodheru ekutarisa. Zvirimwa zvakaiswa pamusoro penzvimbo chena; imwe yakafukidzwa neadjuvant yekurima (Agrotin, Bayer CropScience, Bogotá, Colombia) kuti ifananidze kuvhurwa kwese kwestomata [wet mode (Twet)], uye imwe yaive shizha risina kushandiswa [Dry mode (Tdry)] (Castro -Duque et al., 2020). Kureba pakati pekamera nehari panguva yekutorwa mifananidzo kwaive 1 m.
Relative tolerance index yakaverengerwa zvisina kunanga uchishandisa stomatal conductance (gs) yemiti yakarapwa zvichienzaniswa nemiti yekudzora (miti isina mishonga yekushushikana uye ine zvinodzora kukura zvakashandiswa) kuti ione kuti genotypes dzakarapwa dzakaongororwa muchidzidzo ichi dzinokwanisa kushivirira sei. RTI yakawanikwa uchishandisa equation yakatorwa kubva kuna Chávez-Arias et al. (2020).
Mukuedza kwega kwega, zvese zvinoshanduka mumuviri zvataurwa pamusoro apa zvakaongororwa uye zvakanyorwa pa55 DAE uchishandisa mashizha akakura zvakakwana akaunganidzwa kubva pamusoro pedenga. Pamusoro pezvo, kuyerwa kwakaitwa muimba yekukura kudzivirira kushandura mamiriro ekunze ayo zvirimwa zvinokura.
Data kubva mukuyedza kwekutanga nekwechipiri yakaongororwa pamwe chete senhevedzano yekuedza. Boka rega rega rekuyedza raive nemiti mishanu, uye chirimwa chimwe nechimwe chaive chikamu chekuedza. Kuongororwa kwevariance (ANOVA) kwakaitwa (P ≤ 0.05). Pakaonekwa misiyano yakakosha, bvunzo yekuenzanisa yaTukey post hoc yakashandiswa paP ≤ 0.05. Shandisa basa re arcsine kushandura ma percentage values. Data rakaongororwa uchishandisa Statistix v 9.0 software (Analytical Software, Tallahassee, FL, USA) uye rakarongwa uchishandisa SigmaPlot (vhezheni 10.0; Systat Software, San Jose, CA, USA). Kuongororwa kukuru kwezvikamu kwakaitwa uchishandisa InfoStat 2016 software (Analysis Software, National University of Cordoba, Argentina) kuti vaone varairidzi vekukura kwezvirimwa vakanakisisa vari kudzidza.
Tafura 1 inopfupikisa ANOVA inoratidza kuyedza, marapirwo akasiyana, uye kudyidzana kwadzo nemashizha ephotosynthetic pigments (chlorophyll a, b, total, uye carotenoids), malondialdehyde (MDA) uye proline content, uye stomatal conductance. Mhedzisiro ye gs, relative water content. (RWC), chlorophyll content, chlorophyll alpha fluorescence parameters, crown temperature (PCT) (°C), crop stress index (CSI) uye relative tolerance index yemiti yemupunga pa55 DAE.
Tafura 1. Pfupiso yedata reANOVA pamusoro pezvinhu zvinoshanduka pakugadzira mupunga uye zvemakemikari pakati pekuyedza (ma genotypes) uye kurapwa kwekumanikidzwa kwekupisa.
Musiyano (P≤0.01) mukushandana kwemashizha ane ruvara rwephotosynthetic pigment, huwandu hwechlorophyll (kuverenga kweAtleaf), uye alpha-chlorophyll fluorescence parameters pakati pekuyedza nekurapa zvinoratidzwa muTafura 2. Kupisa kwakanyanya masikati nehusiku kwakawedzera huwandu hwechlorophyll necarotenoid. Mbeu dzemupunga dzisina kupfapfaidzwa kwemashizha ephytohormones (2.36 mg g-1 ye "F67″ uye 2.56 mg g-1 ye "F2000″)) zvichienzaniswa nezvirimwa zvakadyarwa pasi pemamiriro ekunze ekupisa (2.67 mg g -1)) dzakaratidza huwandu hwakaderera hwechlorophyll. Mukuyedza kwese, "F67" yaive 2.80 mg g-1 uye "F2000" yaive 2.80 mg g-1. Pamusoro pezvo, mbesa dzemupunga dzakarapwa nemusanganiswa weAUX neGA sprays pasi pekupisa kwakaratidzawo kudzikira kwehuwandu hwechlorophyll mumarudzi ese ari maviri (AUX = 1.96 mg g-1 uye GA = 1.45 mg g-1 ye "F67"; AUX = 1.96 mg g-1 uye GA = 1.45 mg g-1 ye "F67″; AUX = 2.24 mg) g-1 uye GA = 1.43 mg g-1 (ye "F2000″) pasi pekupisa kwakamanikidzwa. Pasi pekupisa kwakamanikidzwa, kurapwa kwemashizha neBR kwakakonzera kuwedzera kudiki kwekuchinja uku mumarudzi ese ari maviri. Pakupedzisira, CK foliar spray yakaratidza huwandu hwepamusoro hwe photosynthetic pigment pakati pemishonga yese (AUX, GA, BR, SC uye AC treatments) mumarudzi e genotypes F67 (3.24 mg g-1) uye F2000 (3.65 mg g-1). Huwandu hwechlorophyll (Atleaf unit) hwakaderedzwawo nekumanikidzwa kwekupisa kwakabatana. Huwandu hwepamusoro hwakanyorwawo mumiti yakapfapfaidzwa neCC mumarudzi ese ari maviri (41.66 ye "F67" uye 49.30 ye "F2000"). Fv neFv/Fm ratios zvakaratidza mutsauko mukuru pakati pekurapa nemiti yemhando dzakasiyana (Tafura 2). Kazhinji, pakati pezvinhu izvi, F67 yemhando dzakasiyana yakanga isinganyanyi kukanganiswa nekupisa kupfuura F2000 yemhando dzakasiyana. Huwandu hweFv neFv/Fm hwakanyanya kuderera mukuyedza kwechipiri. Mbeu dzakasimba dze 'F2000′ dzisina kupfapfaidzwa ne phytohormones dzaive neFv values yakaderera (2120.15) uye Fv/Fm ratios (0.59), asi kupfapfaidzwa nemashizha emiti neCK kwakabatsira kudzoreredza hunhu uhwu (Fv: 2591, 89, Fv/Fm ratios: 0.73). , kugamuchira kuverenga kwakafanana nekwakanyorwa pamiti ye "F2000" yakadyarwa pasi pemamiriro ekunze ekupisa (Fv: 2955.35, Fv/Fm ratio: 0.73:0.72). Pakanga pasina mutsauko wakakosha mukutanga kwefluorescence (F0), maximum fluorescence (Fm), maximum photochemical quantum yield yePSII (Fv/F0) uye Fm/F0 ratio. Pakupedzisira, BR yakaratidza maitiro akafanana sezvakaonekwa neCK (Fv 2545.06, Fv/Fm ratio 0.73).
Tafura 2. Mhedzisiro yekumanikidzwa kwekupisa kwakabatana (40°/30°C masikati/husiku) pamashizha ane ma "photosynthetic pigments" [total chlorophyll (Chl Total), chlorophyll a (Chl a), chlorophyll b (Chl b) uye carotenoids Cx+c] effect ], relative chlorophyll content (Atliff unit), chlorophyll fluorescence parameters (initial fluorescence (F0), maximum fluorescence (Fm), variable fluorescence (Fv), maximum PSII efficiency (Fv/Fm), photochemical maximum quantum yield yePSII (Fv/F0 ) uye Fm/F0 mumiti yemhando mbiri dzemupunga [Federrose 67 (F67) naFederrose 2000 (F2000)] mazuva makumi mashanu nemashanu mushure mekubuda (DAE)).
Kuwanda kwemvura (RWC) yemiti yemupunga yakarungwa zvakasiyana kwakaratidza mutsauko (P ≤ 0.05) mukudyidzana pakati pemishonga yekuedza neyakurapwa kwemashizha (Mufananidzo 1A). Pakurapwa neSA, kukosha kwakaderera kwakanyorwa kune ese ma genotypes (74.01% yeF67 uye 76.6% yeF2000). Pasi pemamiriro ekumanikidzwa nekupisa, RWC yemiti yemupunga yemarudzi ese maviri akarapwa ne phytohormones akasiyana yakawedzera zvakanyanya. Kazhinji, kushandiswa kweCK, GA, AUX, kana BR pamashizha kwakawedzera RWC kusvika pamitengo yakafanana neyemiti yakakurira mumamiriro ezvinhu akanaka panguva yekuedza. Kudzora kwakakwana uye miti yakapfapfaidzwa pamashizha yakanyora kukosha kweinenge 83% kune ese ma genotypes. Kune rumwe rutivi, gs yakaratidzawo mutsauko wakakosha (P ≤ 0.01) mukudyidzana kwekuyedza nekurapa (Mufananidzo 1B). Chirimwa cheAutified Control (AC) chakanyorawo kukosha kwepamusoro-soro kwegenotype yega yega (440.65 mmol m-2s-1 yeF67 uye 511.02 mmol m-2s-1 yeF2000). Miriwo yemupunga yakapinzwa mukupisa kwakabatana chete yakaratidza kukosha kwakaderera kwe gs kune ese ma genotypes (150.60 mmol m-2s-1 yeF67 uye 171.32 mmol m-2s-1 yeF2000). Kurapa kwemashizha nemidziyo yese inodzora kukura kwezvirimwa kwakawedzerawo g. Pamiti yemupunga yeF2000 yakapfapfaidzwa neCC, mhedzisiro yekupfapfaidza mashizha ne phytohormones yaive pachena. Boka remiti iri harina kuratidza mutsauko kana tichienzanisa nemiti inodzora zvakakwana (AC 511.02 uye CC 499.25 mmol m-2s-1).
Mufananidzo 1. Mhedzisiro yekumanikidzwa kwekupisa pamwe chete (40°/30°C masikati/husiku) pane huwandu hwemvura (RWC) (A), stomatal conductance (gs) (B), malondialdehyde (MDA) kugadzirwa (C), uye huwandu hweproline . (D) muzvirimwa zvema genotype maviri emupunga (F67 naF2000) pamazuva makumi mashanu nemashanu mushure mekubuda (DAE). Kurapa kwakaongororwa kwegenotype yega yega kwaisanganisira: absolute control (AC), heat stress control (SC), heat stress + auxin (AUX), heat stress + gibberellin (GA), heat stress + cell mitogen (CK), uye heat stress + brassinosteroid. (BR). Koramu yega yega inomiririra mean ± standard error yemapoinzi mashanu edata (n = 5). Makoramu anoteverwa nemabhii akasiyana anoratidza mutsauko wakakosha wehuwandu zvichienderana nebvunzo yaTukey (P ≤ 0.05). Mabhii ane chiratidzo chakaenzana anoratidza kuti mean haina kukosha muhuwandu (≤ 0.05).
Zviri muMDA (P ≤ 0.01) uye proline (P ≤ 0.01) zvakaratidzawo mutsauko mukuru mukudyidzana pakati pekuyedza nekurapa nephytohormone (Mufananidzo 1C, D). Kuwedzera kwemafuta eperoxidation kwakaonekwa nekurapwa kweSC mumarudzi ese ari maviri (Mufananidzo 1C), zvisinei zvirimwa zvakarapwa nemashizha ekukura kwemashizha zvakaratidza kudzikira kwemafuta eperoxidation mumarudzi ese ari maviri; Kazhinji, kushandiswa kwephytohormones (CA, AUC, BR kana GA) kunotungamira kudzikira kwemafuta eperoxidation (MDA content). Hapana musiyano wakawanikwa pakati pezvirimwa zveAC zvema genotype maviri nezvirimwa zviri pasi pekupisa uye zvakapfapfaidzwa ne phytohormones (zvakaonekwa kuti FW values muzvirimwa zve "F67" zvakabva pa4.38–6.77 µmol g-1, uye muzvirimwa zve "F2000" "zvakaonekwa kuti values dzakabva pa2.84 kusvika 9.18 µmol g-1 (zvirimwa). Kune rumwe rutivi, kugadzirwa kweproline muzvirimwa zve "F67" kwaive kwakaderera pane muzvirimwa zve "F2000" zviri pasi pekupisa kwakabatana, izvo zvakaita kuti pave nekuwedzera kwekugadzirwa kweproline. muzvirimwa zvemupunga zvine kupisa, mukuyedza kwese, zvakaonekwa kuti kupihwa kwemahormone aya kwakawedzera zvakanyanya amino acid yezvirimwa zveF2000 (AUX neBR zvaive 30.44 uye 18.34 µmol g-1) zvichiteerana (Mufananidzo 1G).
Mhedzisiro yefoliar plant growth regulator spray pamwe nekumanikidzwa kwekupisa pamwe chete pakupisa kwemuti we plant canopy uye relative tolerance index (RTI) zvinoratidzwa muMifananidzo 2A neB. Kune ese ari maviri genotypes, tembiricha yemuti we AC yaive pedyo ne27°C, uye yemiti ye SC yaive inenge 28°C. WITH. Zvakaonekwawo kuti kurapwa kwemuti we foliar ne CK ne BR kwakaita kuti tembiricha yemuti we canopy iderere ne 2–3°C zvichienzaniswa nemiti ye SC (Mifananidzo 2A). RTI yakaratidza maitiro akafanana nedzimwe shanduko dzemuviri, ichiratidza mutsauko mukuru (P ≤ 0.01) mukudyidzana pakati pekuyedza nekurapa (Mifananidzo 2B). Miti ye SC yakaratidza kushivirira kwemuti we genotypes dzese dziri mbiri (34.18% uye 33.52% yemiti yemupunga ye "F67" ne "F2000", zvichiteerana). Kudyisa phytohormones pamashizha kunovandudza RTI mumiti yakatariswa nekupisa kwakanyanya. Mhedzisiro iyi yakanyanya kuoneka mumiti ye "F2000" yakapfapfaidzwa neCC, umo RTI yaive 97.69. Kune rumwe rutivi, misiyano mikuru yakaonekwa chete muyield stress index (CSI) yemiti yemupunga pasi pemamiriro ekumanikidzwa kwefoliar factor (P ≤ 0.01) (Mufananidzo 2B). Miti yemupunga chete ndiyo yakasangana nekumanikidzwa kwekupisa kwakanyanya ndiyo yakaratidza kukosha kwepamusoro kwestress index (0.816). Miti yemupunga payakapfapfaidzwa ne phytohormones dzakasiyana-siyana, stress index yaive yakaderera (values kubva pa0.6 kusvika pa0.67). Pakupedzisira, muti wemupunga wakakura mumamiriro ezvinhu akanaka waive nekukosha kwe0.138.
Mufananidzo 2. Mhedzisiro yekumanikidzwa kwekupisa kwakabatana (40°/30°C masikati/husiku) pakupisa kwedenga (A), relative tolerance index (RTI) (B), uye crop stress index (CSI) (C) yemhando mbiri dzezvirimwa. Mhando dzemupunga wekutengesa (F67 naF2000) dzakarapwa zvakasiyana-siyana zvekupisa. Kurapa kwakaongororwa kwemhando yega yega yemupunga kwaisanganisira: absolute control (AC), heat stress control (SC), heat stress + auxin (AUX), heat stress + gibberellin (GA), heat stress + cell mitogen (CK), uye heat stress + brassinosteroid. (BR). Kumanikidzwa kwekupisa kwakabatana kunosanganisira kuisa zvirimwa zvemupunga pakupisa kwakanyanya kwemasikati/husiku (40°/30°C masikati/husiku). Koramu yega yega inomiririra avhareji ± standard error yemapoinzi mashanu edata (n = 5). Makoramu anoteverwa nemabhii akasiyana anoratidza mutsauko wakakosha wehuwandu zvichienderana nebvunzo yaTukey (P ≤ 0.05). Mabhii ane chiratidzo chakaenzana anoratidza kuti avhareji haina kukosha muhuwandu (≤ 0.05).
Kuongororwa kwezvikamu zvikuru (PCA) kwakaratidza kuti zvinhu zvakaongororwa pa55 DAE zvakatsanangura 66.1% yemhinduro dzemuviri uye dzemakemikari emiti yemupunga yakamanikidzwa nekupisa yakarapwa ne spray inodzora kukura (Mufananidzo 3). Mavectors anomiririra zvinhu uye madots anomiririra zvinodzora kukura kwemiti (GRs). Mavectors e gs, chlorophyll content, maximum quantum efficiency yePSII (Fv/Fm) uye biochemical parameters (TChl, MDA uye proline) ari padivi pekwakabva, zvichiratidza hukama hwakakura pakati pemaitiro emuviri emiti uye izvo. zvinoshanduka. Boka rimwe (V) raisanganisira mbesa dzemupunga dzakarimwa pakupisa kwakanyanya (AT) uye mbesa dzeF2000 dzakarapwa neCK neBA. Panguva imwe chete, mbesa zhinji dzakarapwa neGR dzakaumba boka rakasiyana (IV), uye kurapwa neGA muF2000 kwakaumba boka rakasiyana (II). Kusiyana neizvi, mbesa dzemupunga dzakanyanya kupisa (mapoka I neIII) dzisina ma phytohormones (marudzi ese e genotypes aive SC) dzakawanikwa munzvimbo yakatarisana neboka V, zvichiratidza mhedzisiro yekunyanya kupisa pa physiology yemiti.
Mufananidzo 3. Kuongororwa kwenhoroondo yemhedzisiro yekumanikidzwa kwekupisa kwakabatana (40°/30°C masikati/husiku) pazvirimwa zvemhando mbiri dzemupunga (F67 naF2000) pamazuva makumi mashanu nemashanu mushure mekubuda (DAE). Zvidimbu: AC F67, kutonga kwakazara F67; SC F67, kudzora kupisa F67; AUX F67, kumanikidzwa kwekupisa + auxin F67; GA F67, kumanikidzwa kwekupisa + gibberellin F67; CK F67, kumanikidzwa kwekupisa + kupatsanurwa kwemasero BR F67, kumanikidzwa kwekupisa + brassinosteroid. F67; AC F2000, kutonga kwakazara F2000; SC F2000, Kudzora kupisa F2000; AUX F2000, kumanikidzwa kwekupisa + auxin F2000; GA F2000, kumanikidzwa kwekupisa + gibberellin F2000; CK F2000, kushushikana kwekupisa + cytokinin, BR F2000, kushushikana kwekupisa + steroid yemhangura; F2000.
Zvinhu zvakaita se chlorophyll content, stomatal conductance, Fv/Fm ratio, CSI, MDA, RTI uye proline content zvinogona kubatsira kunzwisisa kugadziriswa kwemhando dzemupunga uye kuongorora kukanganiswa kwemaitiro ekurima pasi pekumanikidzwa kwekupisa (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Chinangwa chekuedza uku kwaive kuongorora mhedzisiro yekushandiswa kwezvinodzora kukura zvina pamatanho emuviri uye emakemikari embeu dzemupunga pasi pemamiriro akaoma ekupisa. Kuedza mbeu inzira iri nyore uye inokurumidza yekuongorora zvirimwa zvemupunga panguva imwe chete zvichienderana nehukuru kana mamiriro ezvivakwa zviripo (Sarsu et al. 2018). Mhedzisiro yechidzidzo ichi yakaratidza kuti kumanikidzwa kwekupisa kwakabatana kunokonzera mhinduro dzakasiyana dzemuviri uye dzemakemikari mumhando mbiri dzemupunga, zvichiratidza maitiro ekugadzirisa. Mhedzisiro iyi inoratidzawo kuti maspray anodzora kukura kwemashizha (kunyanya ma cytokinins nema brassinosteroids) anobatsira mupunga kuti ukwanise kugadzikana nekuomarara kwekupisa sezvo favor inonyanya kukanganisa gs, RWC, Fv/Fm ratio, photosynthetic pigments uye proline content.
Kushandiswa kwezvinodzora kukura kunobatsira kuvandudza mamiriro emvura emiti yemupunga iri pasi pekupisa, izvo zvinogona kunge zvakabatana nekumanikidzwa kwakanyanya uye kudzikira kwekupisa kwemiti. Chidzidzo ichi chakaratidza kuti pakati pemiti yemupunga inonzi "F2000" (inogona kubatwa negenotype), miti yemupunga yakarapwa zvakanyanya neCK kana BR yaive nemazinga e gs akakwira uye PCT yakaderera pane miti yakarapwa neSC. Zvidzidzo zvekare zvakaratidzawo kuti gs nePCT zviratidzo zvemuviri zvakarurama zvinogona kuona mhinduro inochinjika yemiti yemupunga uye mhedzisiro yemaitiro ekurima pakumanikidzwa kwekupisa (Restrepo-Diaz naGarces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). Leaf CK kana BR inowedzera g pasi pekumanikidzwa nekuti mahormone aya emiti anogona kukurudzira kuvhurika kwemupunga kuburikidza nekudyidzana kwekugadzira nemamwe mamorekuru ezviratidzo akadai seABA (promoter of stomatal closure under abiotic stress) (Macková et al., 2013; Zhou et al., 2013). 2013). , 2014). Kuvhurika kwedumbu kunobatsira kutonhora kwemashizha uye kunobatsira kuderedza kupisa kwedenga (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Nekuda kwezvikonzero izvi, tembiricha yedenga remupunga inopfapfaidzwa neCK kana BR inogona kunge yakaderera kana paine kupisa kwakabatana.
Kushushikana zvakanyanya pakupisa kunogona kuderedza huwandu hwepigment yemashizha inogadzirwa nephotosynthesis (Chen et al., 2017; Ahammed et al., 2018). Muchidzidzo ichi, apo zvirimwa zvemupunga zvaive pasi pekupisa kwakanyanya uye zvisina kupfapfaidzwa nezvinogadzirisa kukura kwezvirimwa, pigment yephotosynthesis yaiwanzoderera mumarudzi ese ari maviri (Tafura 2). Feng et al. (2013) vakataurawo kudzikira kukuru kwehuwandu hwechlorophyll mumashizha emarudzi maviri egorosi anoonekwa pasi pekupisa kwakanyanya. Kusangana nekupisa kwakanyanya kunowanzo kukonzera kudzikira kwehuwandu hwechlorophyll, izvo zvinogona kunge zvichikonzerwa nekudzikira kwe chlorophyll biosynthesis, kudzikira kwemavara, kana mhedzisiro yavo yakabatana pasi pekupisa kukuru (Fahad et al., 2017). Zvisinei, zvirimwa zvemupunga zvakarapwa zvakanyanya neCK neBA zvakawedzera huwandu hwemashizha anogadzirwa nephotosynthesis pasi pekupisa kukuru. Mhedzisiro yakafanana yakataurwawo naJespersen naHuang (2015) naSuchsagunpanit et al. (2015), avo vakaona kuwedzera kwehuwandu hwe chlorophyll mumashizha mushure mekushandiswa kwe zeatin ne epibrassinosteroid hormones mu heat-stressed bentgrass nemupunga, zvichiteerana. Tsananguro ine musoro yekuti nei CK ne BR zvichikurudzira kuwedzera kwehuwandu hwe chlorophyll mumashizha pasi pekupisa kwakabatana ndeyekuti CK inogona kuwedzera kutanga kwekusimudzira kwenguva refu kwe expression promoters (senge senescence-activating promoter (SAG12) kana HSP18 promoter) uye kuderedza kurasikirwa kwe chlorophyll mumashizha. , kunonoka kupera kwemashizha uye kuwedzera kuramba kwemiti kukupisa (Liu et al., 2020). BR inogona kuchengetedza chlorophyll mumashizha uye kuwedzera huwandu hwe chlorophyll mumashizha nekuita kuti kana kukurudzira kugadzirwa kwema enzymes anobatanidzwa mu chlorophyll biosynthesis pasi pemamiriro ekumanikidzwa (Sharma et al., 2017; Siddiqui et al., 2018). Chekupedzisira, ma phytohormone maviri (CK neBR) anokurudzirawo kuratidzwa kwemapuroteni ekupisa uye anovandudza maitiro akasiyana-siyana ekugadzirisa metabolic, akadai sekuwedzera kwe chlorophyll biosynthesis (Sharma et al., 2017; Liu et al., 2020).
Chlorophyll, fluorescence parameters, inopa nzira inokurumidza uye isingaparadze inogona kuongorora kushivirira kwezvirimwa kana kugadzirisa mamiriro ekushushikana kweabiotic (Chaerle et al. 2007; Kalaji et al. 2017). Maparamita akadai seFv/Fm ratio akashandiswa sezviratidzo zvekuchinjika kwezvirimwa kumamiriro ekushushikana (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Muchidzidzo ichi, zvirimwa zveSC zvakaratidza kukosha kwakaderera kweizvi, kunyanya zvirimwa zvemupunga zve "F2000". Yin et al. (2010) vakawanawo kuti chiyero cheFv/Fm chemashizha emupunga akanyanya kudiridza chakaderera zvakanyanya patembiricha iri pamusoro pe35°C. Sekureva kwaFeng et al. (2013), chiyero cheFv/Fm chakaderera pasi pekupisa chinoratidza kuti chiyero chekutorwa kwesimba rekukurudzira nekushandurwa nePSII reaction centre chakaderera, zvichiratidza kuti PSII reaction centre inoparara pasi pekupisa. Kuona uku kunotibvumira kugumisa kuti kusagadzikana muzvishandiso zve photosynthetic kunonyanya kuoneka mumhando dzinonzwa kurwara (Fedearroz 2000) pane mumhando dzisingapindi kurwara (Fedearroz 67).
Kushandiswa kweCK kana BR kwakawedzera kushanda kwePSII mumamiriro ezvinhu akaomarara ekumanikidzwa nekupisa. Zvakafanana zvakawanikwa naSuchsagunpanit et al. (2015), avo vakaona kuti kushandiswa kweBR kwakawedzera kushanda kwePSII pasi pekumanikidzwa nekupisa mumupunga. Kumar et al. (2020) vakawanawo kuti zvirimwa zvechickpea zvakarapwa neCK (6-benzyladine) uye zvakaiswa pasi pekumanikidzwa nekupisa zvakawedzera chiyero cheFv/Fm, zvichigumisa kuti kushandiswa kweCK pamashizha nekuita kuti zeaxanthin pigment cycle ishande zvakanaka kwakasimudzira PSII. Pamusoro pezvo, BR leaf spray yaifarira PSII photosynthesis pasi pemamiriro ezvinhu ekumanikidzwa, zvichiratidza kuti kushandiswa kwephytohormone iyi kwakakonzera kudzikira kwekupararira kwesimba rekumutsa rePSII antennae uye kwakakurudzira kuunganidzwa kwemapuroteni madiki ekumanikidzwa nekupisa muma chloroplasts (Ogweno et al. 2008; Kothari naLachowitz). , 2021).
Zviri mukati meMDA neproline zvinowanzowedzera kana zvirimwa zviri pasi pekumanikidzwa kweabiotic zvichienzaniswa nezvirimwa zvinorimwa mumamiriro ezvinhu akanaka (Alvarado-Sanabria et al. 2017). Zvidzidzo zvekare zvakaratidzawo kuti MDA neproline levels zviratidzo zvebiochemical zvinogona kushandiswa kunzwisisa maitiro ekuchinja kana kukanganisa kwemaitiro ekurima mupunga pasi pekupisa kwakanyanya kwemasikati kana husiku (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al. . , 2021). Zvidzidzo izvi zvakaratidzawo kuti MDA neproline zviri mukati zvaiwanzova zvakakwirira mumiti yemupunga inoratidzirwa kukupisa kwakanyanya husiku kana masikati, zvichiteerana. Zvisinei, kupfapfaidza CK neBR pamashizha kwakabatsira kudzikira kweMDA uye kuwedzera kwemazinga eproline, kunyanya mu genotype inoshivirira (Federroz 67). CK spray inogona kukurudzira kuwanda kwe cytokinin oxidase/dehydrogenase, nokudaro ichiwedzera huwandu hwemakemikari anodzivirira akadai se betaine neproline (Liu et al., 2020). BR inokurudzira kupinza ma osmoprotectorants akadai se betaine, shuga, uye amino acids (kusanganisira free proline), ichichengetedza masero e osmotic balance pasi pemamiriro ezvinhu akawanda asina kunaka (Kothari naLachowiec, 2021).
Crop stress index (CSI) uye relative tolerance index (RTI) zvinoshandiswa kuona kana kurapwa kuri kuongororwa kuchibatsira kuderedza kushushikana kwakasiyana-siyana (kusashanda uye kusashanda) uye kuva nemigumisiro yakanaka pakushanda kwemiti (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). CSI values inogona kubva pa0 kusvika pa1, zvichimiririra mamiriro asina kushushikana uye kusashanda, zvichiteerana (Lee et al., 2010). CSI values yemiti ine kupisa (SC) yakabva pa0.8 kusvika pa0.9 (Mufananidzo 2B), zvichiratidza kuti miti yemupunga yakakanganiswa zvakanyanya nekumanikidzwa kwakabatana. Zvisinei, kupfapfaidza BC (0.6) kana CK (0.6) zvakanyanya kukonzera kudzikira kwechiratidzo ichi pasi pemamiriro ekumanikidzwa kwemiti zvichienzaniswa nemiti yemupunga yeSC. Muzvirimwa zveF2000, RTI yakaratidza kuwedzera kwakanyanya pakushandisa CA (97.69%) uye BC (60.73%) zvichienzaniswa neSA (33.52%), zvichiratidza kuti izvi zvinodzora kukura kwezvirimwa zvinobatsirawo mukuvandudza mhinduro yemupunga pakushivirira kwemusanganiswa. Kupisa zvakanyanya. Aya maindices akakurudzirwa kugadzirisa mamiriro ekushushikana mumarudzi akasiyana. Chidzidzo chakaitwa naLee et al. (2010) chakaratidza kuti CSI yemhando mbiri dzedonje pasi pekumanikidzwa kwemvura zvine mwero yaive inenge 0.85, nepo CSI kukosha kwemhando dzakadiridzwa zvakanaka kwaive kubva pa0.4 kusvika pa0.6, zvichigumisa kuti iyi index chiratidzo chekuchinja kwemvura kwemhando. mamiriro ekushushikana. Uyezve, Chavez-Arias et al. (2020) vakaongorora kushanda kwezvinogadzirisa zvakagadzirwa senzira yakazara yekugadzirisa kushushikana muzvirimwa zveC. elegans uye vakawana kuti zvirimwa zvakapfapfaidzwa nemishonga iyi zvakaratidza RTI yakakwira (65%). Zvichibva pane zviri pamusoro apa, CK neBR zvinogona kutariswa senzira dzekurima dzakanangana nekuwedzera kushivirira kwemupunga kukupisa kwakaoma, sezvo izvi zvinodzora kukura kwezvirimwa zvichikonzera mhinduro dzakanaka dzemakemikari nemuviri.
Mumakore mashoma apfuura, tsvakurudzo yemupunga muColombia yakatarisana nekuongorora majini anotsungirira tembiricha yepamusoro yemasikati kana husiku uchishandisa hunhu hwemuviri kana hwemakemikari (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Zvisinei, mumakore mashoma apfuura, ongororo yematekinoroji anoshanda, anochengetedza mari uye anobatsira yave kukosha zvikuru kupa mazano ekutarisira zvirimwa zvakabatana kuti zvivandudze mhedzisiro yenguva dzakaoma dzekushushikana kwekupisa munyika (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021). Saka, mhinduro dzemuviri uye dzemakemikari emiti yemupunga kukutambudzika kwakaoma (40°C masikati/30°C husiku) dzakaonekwa muchidzidzo ichi dzinoratidza kuti kupfapfaidza mashizha neCK kana BR kungave nzira yakakodzera yekutarisira zvirimwa kuderedza migumisiro yakaipa. Mhedzisiro yenguva dzekushushikana kwekupisa zvine mwero. Kurapa uku kwakavandudza kushivirira kwemarudzi ese emupunga (low CSI neRTI yakakwira), zvichiratidza maitiro akajairika mumhinduro dzemuviri uye dzemakemikari emiti pasi pekumanikidzwa kwekupisa kwakabatana. Mhinduro huru yezvirimwa zvemupunga yaive kudzikira kwehuwandu hweGC, huwandu hwechlorophyll, chlorophylls α uye β uye carotenoids. Pamusoro pezvo, zvirimwa zvinotambura nekukuvadzwa kwePSII (kuderera kwe chlorophyll fluorescence parameters dzakadai seFv/Fm ratio) uye kuwedzera kwe lipid peroxidation. Kune rumwe rutivi, apo mupunga wakarapwa neCK neBR, mhedzisiro iyi yakaipa yakaderedzwa uye huwandu hweproline hwakawedzera (Mufananidzo 4).
Mufananidzo 4. Muenzaniso wepfungwa wemhedzisiro yekumanikidzwa kwekupisa pamwe chete uye mushonga wekukura kwezvirimwa pamiti yemupunga. Miseve mitsvuku nebhuruu inoratidza mhedzisiro yakaipa kana yakanaka yekudyidzana pakati pekumanikidzwa kwekupisa uye kushandiswa kweBR (brassinosteroid) neCK (cytokinin) pamhinduro dzemuviri nedzemakemikari, zvichiteerana. gs: stomatal conductance; Total Chl: huwandu hwechlorophyll; Chl α: huwandu hwechlorophyll β; Cx+c: huwandu hwecarotenoid;
Muchidimbu, mhinduro dzemuviri uye dzemakemikari muchidzidzo ichi dzinoratidza kuti zvirimwa zvemupunga zveFedearroz 2000 zviri nyore kubatwa nenguva yekumanikidzwa kwakaoma kwekupisa kupfuura zvirimwa zvemupunga zveFedearroz 67. Vese vanodzora kukura vakaongororwa muchidzidzo ichi (auxins, gibberellins, cytokinins, kana brassinosteroids) vakaratidza kudzikiswa kwekupisa kwakabatana. Zvisinei, cytokinin nebrassinosteroids zvakakonzera kugadziriswa kuri nani kwezvirimwa sezvo zvese zvinodzora kukura kwezvirimwa zvakawedzera huwandu hwechlorophyll, alpha-chlorophyll fluorescence parameters, gs neRWC zvichienzaniswa nezvirimwa zvemupunga zvisina kushandiswa, uye zvakare kudzikiswa kwehuwandu hweMDA nekupisa kwedenga. Muchidimbu, tinogumisa kuti kushandiswa kwezvinodzora kukura kwezvirimwa (cytokinins nebrassinosteroids) chishandiso chinobatsira mukugadzirisa mamiriro ekumanikidzwa muzvirimwa zvemupunga zvinokonzerwa nekumanikidzwa kwakanyanya kwekupisa panguva dzekupisa kwakanyanya.
Zvinyorwa zvepakutanga zvakaratidzwa muchidzidzo ichi zvakabatanidzwa munyaya yacho, uye mimwe mibvunzo inogona kuendeswa kumunyori anoenderana nazvo.
Nguva yekutumira: Nyamavhuvhu-08-2024